Asymmetric habitat isolation and sexual isolation predicted by the cost of migration and hybridization generate novel signatures of reinforcing selection

While ecology and geography can play important roles in the evolution of reproductive isolation across the speciation continuum, the few studies to date examining their relative contributions have largely focused on the early stages of speciation. Studies exploring the evolution of multiple reproductive barriers among divergent lineages, where speciation is nearly complete, are also needed to gain a fuller understanding of the mechanisms promoting and constraining the speciation process. We examine the later stage of speciation by comparing the strength of habitat isolation and sexual isolation among closely related species of gall-forming wasps in the genus Belonocnema experiencing divergent ecologies due to different hos plant use and variable rates of migration due to geography. We found that the strength of both habitat isolation and sexual isolation is lowest among allopatric species pairs with the same host plant association, followed by allopatric species with different host plant association, and highest between sympatric species with different host-plant associations. This pattern strongly suggests that both ecology due to divergent host use, and geography contribute to the evolution of these two reproductive barriers. Notably, reproductive character displacement contributes to nearly half of the strength of both habitat isolation and sexual isolation in sympatry.